Evolutionary Genetics in the Wild : from Populations to Individuals

Predicting evolutionary outcomes and reconstructing past evolutionary transitions are among the main goals of evolutionary biology. Ultimately, understanding the mechanisms of evolutionary change will also provide answers to the timely question of whether and how organisms will adapt to changing environmental conditions. In this thesis, I have investigated the relative roles of natural selection, random genetic drift and genetic correlations in the evolution of complex traits at different levels of organisation from populations to individuals. I have shown that natural selection has been the driving force behind body shape divergence of marine and freshwater threespine stickleback (Gasterosteus aculeatus) populations, while genetic drift may have played a significant role in the more fine scale divergence among isolated freshwater populations. These results are concurrent with the patterns that have emerged in the published studies comparing the relative importance of natural selection and genetic drift as explanations for population divergence in different traits and taxa. I have also shown that body shape and armour divergence among threespine stickleback populations is likely to be biased by the patterns of genetic variation and covariation. Body shape and armour variation along the most likely direction of evolution the direction of maximum genetic variance reflects the general patterns of variation observed wild populations across the distribution range of the threespine stickleback. Conversely, it appears that genetic correlations between the sexes have not imposed significant constraints on the evolution of sexual dimorphism in threespine stickleback body shape and armour. I have demonstrated that the patterns of evolution seen in the wild can be experimentally recreated to tease out the effects of different selection agents in detail. In addition, I have shown how important it is to take into account the correlative nature of traits, when making interpretations about the effects of natural selection on individual traits. Overall, this thesis provides a demonstration of how considering the relative roles of different mechanism of evolutionary change at different levels of organisation can aid in an emergence of a comprehensive picture of how adaptive divergence in wild populations occurs.Evoluutiobiologian keskeisimpiä tavoitteita on lajien evolutiivisen historian selvittäminen ja tulevaisuudessa tapahtuvan evoluution ennustaminen. Näiden tavoitteiden saavuttamisen tarkoitus on puolestaan antaa vastaus yhteen biologian tärkeimmistä ja ajankohtaisimmista kysymyksistä kuinka organismit sopeutuvat muuttuvaan ympäristöön? Ympäristö määrittelee, minkälaiset luonnonvalintapaineet lajeihin kohdistuvat, ja geeniperimä sen, kuinka voimakkaasti luonnonvalinta voi lajeja muuttaa. Populaatiossa täytyy olla geenien erilaisia muunnelmia, jotta evoluutiota voisi tapahtua. Lajien evoluutiota voi kuitenkin tapahtua ilman luonnonvalintaa, jos jotkut geenien muodot yleistyvät populaatiossa sattumanvaraisesti eli geneettisen ajautumisen seurauksena. Riippumatta evoluution takana olevasta voimasta, yksittäisten ominaisuuksien evoluutio tapahtuu harvoin ilman, että jokin toinen ominaisuus muuttuu samalla. Tähän on syynä se, että ominaisuudet eivät ole toisistaan geneettisesti riippumattomia sama geeni voi esimerkiksi vaikuttaa useampaan ominaisuuteen tai geenit, jotka vaikuttavat eri ominaisuuksiin, ovat perimässä fyysisesti lähekkäin eli ominaisuuksien välillä vallitsee geneettinen korrelaatio. Väitöskirjassani olen tutkinut mekanismeja, jotka vaikuttavat kolmipiikin (Gasterosteus aculeatus) ruumiinmuodon ja suojavarustuksen evoluutioon. Olen pyrkinyt selvittämään, mikä osuus kolmipiikkien evoluutiossa on luonnonvalinnalla, geneettisellä ajautumisella ja geneettisillä korrelaatioilla. Väitöskirjani osoittaa, että luonnonvalinta on ollut voimakkain tekijä meri- ja järvipopulaatioiden välillä tapahtuneeseen erilaistumiseen, kun taas geneettinen ajautuminen on mahdollisesti ollut tärkeä tekijä järvipopulaatioiden välisessä erilaistumisessa. Vastaavat tulokset ovat yleisiä muidenkin lajien ja ominaisuuksien kohdalla. Tämä selviää tekemästäni tilastollisesta analyysistä, jossa on koottu yhteen tulokset tutkimuksista, jotka ovat vertailleet luonnonvalinnan ja geneettisen ajautumisen suhteellisia osuuksia eri lajien ja ominaisuuksien evoluutiossa. Väitöskirjatutkimukseni osoittaa myös, että kolmipiikkien ruumiinmuodon ja suojavarustuksen evoluutioon vaikuttaa myös se, kuinka paljon ja minkälaisia muunnelmia näihin ominaisuuksiin vaikuttavista geeneistä alkuperäisessä populaatiossa on. Myös näiden ominaisuuksien väliset geneettiset korrelaatiot vaikuttavat evoluutioon. Voidaankin olettaa evoluution tapahtuvan suurimmalla todennäköisyydellä niissä ominaisuuksissa, joissa geenien eri muunnelmien määrä on suurin. Geneettiset korrelaatiot on kuitenkin otettava huomioon. Tutkimukseni osoittaa, että evoluution suuntaa luonnonpopulaatioissa voidaan ennustaa tarkastelemalla geenien muunnelmien määrää ja geneettisiä korrelaatioita kantapopulaatiossa. Geneettiset korrelaatiot eivät kuitenkaan täysin rajoita kolmipiikkien evoluutiota, mistä on osoituksena sukupuolien välisten erojen evoluutio myös niissä ominaisuuksissa, joihin vaikuttavien geenien välillä on naaraiden ja koiraiden välillä korrelaatioita. Tutkimukseni todistavat, että luonnossa havaittavia evoluutiomalleja voidaan myös luoda kokeellisesti. Nämä kokeet ovat myös vahvistaneet käsitystä ominaisuuksien välisten geneettisten korrelaatioiden huomioonottamisen tärkeydestä tulkittaessa luonnonvalinnan vaikutuksia yksittäisiin ominaisuuksiin. Kokonaisuutena väitöstutkimukseni havainnollistaa, että tarkastelemalla evoluution vaikuttavien tekijöiden suhteellisia osuuksia eri tasoilla, voidaan saada kokonaisvaltainen kuva luonnonvaraisten populaatioiden sopeutumisesta muuttuvaan ympäristöön

Keilaavan millimetriaaltoradiolinkin suuntaaminen ja seuraaminen

In order to provide high-throughput mobile broadband in a dense urban information society, upcoming cellular networks will finally employ the under-utilized millimeter-wave (mmW) frequencies. The challenging mmW radio environment, however, necessitates massive cell densification with wireless backhauling using very directional links. This thesis investigates how these links between access points may be aligned efficiently, and how alignment reflects the network organization. The work provides a thorough presentation of different high-level aspects and background information required when designing a mmW small cell system. In terms of alignment functionality, both automatic link establishment and proactive tracking are considered. Additionally, the presentation includes an overview of beam steerable antennas, mmW propagation in urban environments, and network organization. The thesis further specifies requirements, proposes possible approaches and compares those with existing implementations. Most of existing mmW beam alignment solutions are intended for short-range indoor communications and do not address the issues in cellular systems. While existing functionality considers only a single link between two devices, efficient design should consider both the entire network and the underlying phenomena. The devices should further exploit the existing network infrastructure, location and orientation information, and the concepts of machine learning. Even though the world has recently seen advancements in the related fields, there is still much work to be done before commercial deployment is possible.Seuraavan sukupolven matkaviestinjärjestelmien erittäin nopeissa datayhteyksissä tullaan hyödyntämään millimetriaaltoteknologiaa. Näillä taajuuksilla radioympäristö on kuitenkin hyvin haastava, mikä edellyttää verkon solutiheyden moninkertaistamista, täysin langattomia tukiasemia ja erittäin suuntaavia antenneja. Tässä diplomityössä tutkitaan eri keinoja kuinka tukiasemien väliset linkit kohdistetaan tehokkaasti, ja miten se vaikuttaa verkon rakenteeseen ja hallintaan. Työ tarjoaa kattavan taustaselvityksen mm-aaltosoluverkon toteuttamiseen tarvittavista asioista. Keilanohjausta tarkastellaan sekä verkon automaattisen laajentamisen että kohteen aktiivisen seurauksen kannalta. Tämän lisäksi työssä tutkitaan keilattavia antenneja, mm-aaltojen etenemistä kaupunkiympäristöissä ja verkkorakennetta. Näiden lisäksi työssä rajataan edellytykset, esitetään mahdollisia ratkaisuja, ja vertaillaan näitä olemassa oleviin toteutuksiin. Nykyiset keilaustoteutukset ovat pääasiassa suunniteltu lyhyen kantaman sisäyhteyksille, eivätkä siten vastaa ongelman asettelua. Aikaisempi toiminnallisuus keskittyy yhteen ainoaan linkkiin vaikka tehokas toteutus huomioisi koko järjestelmän kohdistusongelman fysikaalista perustaa unohtamatta. Verkkolaitteiden tulisi hyödyntää olemassa olevaa radioverkkoa, sekä paikka- että suuntatietoja, ja koneoppimisen keinoja. Vaikka aiheeseen liittyvä teknologia on kehittynyt viime vuosina harppauksin, mm-aaltosoluverkot ovat kaikkea muuta kuin valmiita markkinoille

Nordic Migration and Integration Research : Overview and Future Prospects

EXECUTIVE SUMMARY Migration and integration are currently highly contentious topics in political, public and scientific arenas, and will remain so in the near future. However, many common migration-related prejudices and inefficien¬cies in the integration of the migrant population are due to the lack of sound, tested and accessible scientific research. Therefore, the study of migration – by developing basic research and by properly resourcing novel methodological approaches and interventions – will be instrumental in providing a better knowledge base for decision-makers and promoting a more informed population at large. This is the grand purpose this overview report seeks to facilitate. This report was commissioned by NordForsk in August 2016, and has been written by two senior researchers at the Migration Institute of Finland (MIF), Niko Pyrhönen and Johanna Leinonen, with supervision by MIF’s director Tuomas Martikainen. It is based on two main sources: 1) interviews with 56 Nordic experts on migration and integration conducted in Denmark, Finland, Iceland, Norway, and Sweden in late 2016, and 2) an online survey of 356 respondents, distributed to researchers via mailing lists of several Nordic research networks and institutions. The report also includes an overview of current migration trends, a review of central aspects of Nordic migration and integration research, and an overview of research infrastructure in the field. The authors set out on this five-month project with an ambitious goal of charting the current state of Nordic field of migration and integration research. On the one hand, the importance and topicality of this work have been underlined by an emerging sense of global and local migration crisis in the aftermath of a sudden and rapid influx of refugees and asylum-seekers into Europe since the autumn of 2015. On the other hand, an overview such as this is also needed to develop Nordic research collaboration in a contested, polarized and politicized field. We believe that documenting, analyzing and distributing critical assessment and constru¬ctive ideas expressed by researchers with a remarkable array of disciplinary and thematic expertise can help unleash a wide range of unrealized potential and further develop Nordic added value to its fullest extent. The most important results are outlined in this summary, and discussed in more detail in the main body of the report. While all the respondents who participated in this project acknowledge the salience of the crisis sentiment in public, political and scientific debates, most would also like to engage in research that further problema¬tizes and deconstructs the very term ‘refugee crisis’. Considering that the number of international migrants has increased by 60 percent since 1990, and that the number of refugees in 2014 was the highest since World War II, it seems evident that there was a crisis already before the crisis. Demographers, human rights lawyers, and researchers of international relations point out that the rising volume of asylum-seekers has revealed a general disillusionment in the ability of international conventions to adapt properly to ‘the new age of migrations’. As a result, more and more migrants are falling between rigid legal categories and thus being defined as ‘irregular’. However, many respondents point out that the local manifestation of this crisis is often one of national solidarity, which especially in the Nordic context has long been marked by a mythical yearning for a common national purpose exemplified in the Swedish concept of 'folkhemmet', the peoples’ home. The formation of distinct political fault lines between those emphasizing global responsibility of the Global North for the refugees and those advocating the use of advanced border measures to bring migrancy under political control started as far back as the early 2000s. Many respondents are worried about a similar polarization of research into ‘humanist’ and ‘technocratic’ ‘epistemic communities’. Such a development is particularly evident in the endeavors to conceptualize migration as a ‘stress test’ for the Nordic welfare state. There is consensus among the research community that directly policy-relevant approaches are needed to create more resilient systems for integrating migrants. Indeed, excellent register and census data make the Nordic countries a unique comparative context for ‘most similar’ research designs. However, in order to find corroboration between different studies and empirically gauge for best practices, the respondents call for more harmonized data, definitions and framings – supported by theoretical development that challenges methodological nationalism and the notion of Nordic exceptionalism. Even though most Nordic research institutions consist of markedly multidisciplinary staff, there is much room for improvement in cooperation between humanists and social scientists. Respondents agree that rigorous nesting of qualitative and quantitative approaches in the same research projects and work packages is called for in order to reach more sound empirical conclusions. It is worth noting that the researchers themselves are overwhelmingly positive towards increased introduction of such nested approaches. A commonly raised concern, however, is that the funders are not able to properly merit or incentivize experimental, mixed-met¬hod projects, and rarely request explicitly that these projects are to be completed in collaboration between institutions from more than one Nordic country. When asked to discuss specific gaps in the contemporary Nordic research on migration and integration, researchers list methodological questions as being among the most common concerns. Here most of the respondents underline the importance of more careful and expansive historical contextualization and network-embedded research of informal movements. Researchers of civil society and the media sphere hope to see new computational methods adopted to tackle big data-related hurdles, possibly also offering prognostics on how to counter increasing xenophobia. Health and welfare researchers, in particular, point to the need to introduce longitudinal and ‘life-course’ follow-up projects and action research components with scalable intervention pilots. The respondents acknowledge that policy-oriented research on certain topics, such as urban segregation, structural discrimination and large-scale labor migration, has matured much more across the Atlantic. As such, many advocate searching for more appropriate benchmarks for integration success and failure outside the Nordic context – instead of merely measuring employment and income gaps between the minority and majority populations against those in other Nordic countries. Most of the experts interviewed further suggest that focusing on minorities and the autochthonous population simultaneously is a particularly fruitful means for examining, illustrating and developing Nordic added value in migration and integration research. The array of challenges is not limited to difficulties in crossing disciplinary boundaries, but is also reflected in the isolation of themes that are pursued in Nordic research projects. Such segregation can be seen, for example, in how policies and their effects in key research streams – movement, settlement and control measures – are commonly assessed by individuals in different research programs or work packages. While the increased availability of funding for sector-based research, especially in the aftermath of the crisis, is acknowledged as a good thing in principle, it also serves to further fragment the research field into more narrowly focused competencies. As rising research areas – such as forced migration, discrimination and health and well-being – are starting to reach the traction held by areas such as integration, labor market and education, there are even more reasons and more avenues for funders to develop Nordic research by demanding synthesizing, nesting and mixed-method approaches

Kannattaako työpajatoiminta? : Tutkimus työpajojen taloudellisesta ja sosiaalisesta toiminnasta

Tässä lyhennelmässä on tiiviissä muodossa esitetty v. 2001 toteutetun nuorten työpajatoimintaa koskevan tutkimuksen tulokset. Tutkimuksessa tarkasteltiin nuorten työpajatoiminnan kannattavuutta sekä sen taloudellisen tuloksen että sen sosiaalisen vaikuttavuuden avulla. Samalla arvioitiin työpajojen kehittämismahdollisuuksia. Tutkimuksessa työpajoja tarkastellaan sosiaalisina yhteisöinä, jotka toteuttavat yhtäältä sisällöllisiä tavoitteitaan asiakasryhmiensä tukemiseksi ja toisaalta tuottavat tuotteita ja palveluja liiketoimintatavoitteidensa saavuttamiseksi. Tarkastelussa hyödynnetään SYTA-mallia ® (Sosiaalisen Yrityksen / Yhteisön Toiminnan Analysointi), jolla voidaan selvittää organisaation toiminnan taloudellisuutta rahoittajan näkökulmasta. Sosiaalisen tuen vaikuttavuutta ja pajojen toimintaa arvioidaan pajanuoriin, pajojen henkilökuntaan ja niiden ulkoisiin sidosryhmiin kohdistuvien haastattelu- ja kyselyaineistojen perusteella. Sosiaalisella tuella tässä tutkimuksessa tarkoitetaan tietoista ja tavoiteltua prosessia työpajanuoren elämän kokonaisvaltaiseksi tukemiseksi - sekä sosio-emotionaalisen auttamisen että työhön tai koulutukseen liittyvän tuen osalta. Aineiston perusteella voidaan todeta, että rahoittajan (yhteiskunta) kannalta työpajatoimintaan kohdistetut panostukset ovat mielekkäitä siihen käytettyjen todellisten lisäpanosten (toteutuneiden nettokustannusten ja vaihtoehtoisten nettokustannusten erotus) ja saavutettujen sisältötulosten suhteen arvioituna. Arviolta noin puolet työpajanuorista sijoittuu ns. aktiivitoimiin työpajan jälkeen, ja noin kolmasosa palaa työttömyyteen. Työpajatoiminnan vahvuuksia ovat toimiva sisäinen vuoro- vaikutus, jolla viitataan läheisiin ja toimiviin ohjaajakokemuksiin sekä onnistuneeseen sosiaaliseen tukeen. Kehittymisalueena työpajoilla on ulkoisen vuorovaikutuksen eli yhteistyösuhteiden selkiyttäminen alueen muiden toimijoiden kanssa. Kehittämisehdotuksina esitetään, että toiminnan jatkuvuuden turvaamiseksi pajat irrottautuisivat omiksi juridisiksi yksiköikseen ja tuotteistaisivat palvelujaan siten, että projektimuotoisen rahoituksen päättyminen ei ajaisi toimintaa kriisiin. Pajojen kannalta irrottautuminen muodostaisi paineen oman toiminnan kehittämiseen ja toimintaan liittyvän liiketoiminnan ylläpitämiseen. Tämä ei tarkoita sitä, että sosiaalisen yhteisön yleistuen tarve välttämättä poistuisi. Tällä tuella kompensoidaan niitä kustannuksia, joita muodostuu pyrittäessä asiakasryhmien elämänhallinnan tukemiseen. Muutos organisaation kehittämiseksi tapahtuisi selkiyttämällä työpajan perustehtävää (miksi ollaan olemassa), ottamalla liike- toiminta-ajattelua sisältötoiminnan rinnalle (elämän tukeminen + taloudelliset realiteetit ja organisaation kehittäminen) ja avointa yhteistyötä sidosryhmien kanssa harjoittamalla (yritysyhteistyö sekä asiakasryhmien kannalta keskeisten toimijoiden yhteiset prosessit)

Restored river habitat provides a natural spawning area for a critically endangered landlocked Atlantic salmon population

Supplementing endangered fish populations with captive bred individuals is a common practice in conservation management. The aim of supplementary releases from hatchery broodstocks is to maintain the viability of populations by maintaining their genetic diversity. Landlocked Lake Saimaa salmon (Salmo salar m. sebago) has been critically endangered for the past half-century. As a result of anthropogenic disturbance, especially construction of hydroelectric power plants, the Lake Saimaa salmon has become completely dependent on hatchery broodstock. Recently, habitat restoration has been done in one of the former spawning rivers with the aim of creating a new natural spawning ground for the critically endangered population. Hatchery fish releases have also been revised so that in addition to juveniles, adult fish from the hatchery and from the wild have been released into the restored river. We assessed here if a restored river stretch can be used as a natural spawning ground and juvenile production area with the aim of improving genetic diversity of the critically endangered Lake Saimaa salmon. By constructing a pedigree of the released adults, and juveniles sampled from the restored river, we found that the majority of the released adults had produced offspring in the river. We also found that wild-caught spawners that were released into the restored river had much higher reproductive success than hatchery-reared parents that were released into the restored river at the same time. We found no significant differences in genetic diversity between the parent and offspring generations. Meanwhile, relatedness among different groups of adults and juveniles varied a lot. For example, while the hatchery-reared females were on average half-siblings, wild-caught females showed no significant relatedness. This highlights the importance of using pedigree information in planning the conservation and management of endangered populations, especially when artificial propagation is involved.Peer reviewe

Inheritance of Vertebral Number in the Three-Spined Stickleback (Gasterosteus aculeatus)

Peer reviewe

Upper limb dysfunction and activities in daily living in idiopathic normal pressure hydrocephalus

Background Idiopathic normal pressure hydrocephalus (iNPH) is a neurodegenerative disease with a characteristic symptom triad of gait disturbance, cognitive decline, and incontinence. Recently, also dysfunctions in upper limbs have been described in iNPH and reported to improve after shunt surgery. We aim to describe the role of upper limb motor function in the clinical assessment of iNPH patients and its influence on activities of daily living (ADL). Methods Seventy-five consecutive patients with probable iNPH were studied pre-operatively and at 3 and 12 months after shunt surgery. The pre-operative evaluation included lumbar drainage of cerebrospinal fluid (tap test). Motor functions were assessed in upper and lower limbs with Grooved Pegboard Test (GPT), Box & Block Test (BBT), Total Score of Gait (TSG), and balance test. ADL was assessed with Barthel's index and cognition in accordance with the Consortium to Establish a Registry for Alzheimer's Disease (CERAD). Results Patients showed improvement in all motor tests and ADL at 3 months after shunt surgery. The improvement remained stable during the 12-month post-operative follow-up. The motor function tests correlated with each other and with ADL. Conclusions A 3-month follow-up period after shunt surgery is adequate to show improvement in motor tasks, and a positive outcome will last for at least 12 months. A shunt-responsive dysfunction of upper limb motor performance plays a major role in ADL of iNPH patients. Therefore, we suggest an evaluation of upper limb motor performance to be included in routine evaluation of iNPH patients.Peer reviewe

Heritability of Asymmetry and Lateral Plate Number in the Threespine Stickleback

Peer reviewe

Cross-cultural adaptation and validation of the Finnish version of the central sensitization inventory and its relationship with dizziness and postural control

Background: Central Sensitization (CS) involves dysfunction in neurophysiological mechanisms that increase neuronal responses to both noxious and non-noxious stimuli in the central nervous system. The Central Sensitization Inventory (CSI) is considered the leading patient-reported outcome measure for assessing CS-related symptoms. The aim of this study was to translate and cross-culturally adapt the CSI into Finnish (CSI-FI) and to evaluate its psychometric properties. Methods: Translation and cross-cultural validation of the CSI was conducted according to established guidelines. The validation sample was 229 subjects, including 42 pain free controls and 187 subjects with chronic musculoskeletal pain. The CSI-FI was evaluated for internal consistency, test-retest reliability, exploratory factor analysis with maximum likelihood extraction, relationship with subject-reported outcome measures [Tampa scale of kinesiophobia (TSK), the Depression scale (DEPS), 5-level EQ-5D version (EQ-5 L-5D), Roland-Morris Disability Questionnaire (RMDQ), and Pain and Sleep Questionnaire Three-Item Index (PSQ-3)], pain history, subjective symptoms of dizziness, and CS-related diagnoses on CSI part B. Furthermore, we studied the ability of the CSI-FI to distinguish pain free controls, subjects with chronic pain in a single body area, and subjects with multisite chronic pain. In addition, we studied the relationship of CSI-FI scores with postural control on a force plate. Results: The CSI-FI demonstrated good internal consistency (0.884) and excellent test-retest reliability (0.933) with a 7 ± 1 day gap between test administrations. Exploratory factor analysis with maximum likelihood extraction yielded a one factor solution. Fair to good correlations were found between the CSI-FI and the TSK, DEPS, EQ-5 L-5D, RMDQ, and PSQ-3. Subjective symptoms of dizziness correlated better with CSI-FI scores than any of the CS-related diagnoses on CSI part B. Total CSI-FI scores successfully distinguished between pain free controls, subjects with chronic pain in a single body area, and subjects with multisite chronic pain. The multisite pain group reported significantly more dizziness symptoms than the other two groups. Force plate measurements showed no relationship between postural control and CSI-FI scores. Conclusion: The CSI-FI translation was successfully cross-culturally adapted and validated into Finnish. CSI-FI psychometric properties and scores were all in acceptable levels and in line with previous CSI validations. The CSI-FI appears to be a valid and reliable instrument for assessing CS-related symptomology in Finnish-speaking populations